|Distribution of the sand cat in 2016|
The sand cat (Felis margarita), also known as the sand dune cat, is the only cat living chiefly in true deserts. This small cat is widely distributed in the deserts of North Africa, the Middle East and Central Asia. Starting in 2002, it was listed as near threatened on the IUCN Red List because the population was considered fragmented and small with a declining trend. It was downlisted to least concern in 2016.
Owing to long hairs covering the soles of its feet, the sand cat is well adapted to the extremes of a desert environment and tolerant of extremely hot and cold temperatures. It inhabits both sandy and stony deserts, in areas far from water sources.
Felis margarita was the scientific name proposed by Victor Loche in 1858 who first described a sand cat specimen found in the area of "Négonça" in the northern Algerian Sahara. This holotype specimen appears to have been lost. The species was named after the French General Jean Auguste Margueritte. In the 20th century, the following zoological specimens of sand cats were described:
- Eremaelurus thinobius was proposed as a species by Sergey Ognev in 1926. This specimen had been collected in the Karakum Desert in Turkmenistan. In 1938, Reginald Innes Pocock also considered it a species, but subordinated it to the genus Felis using the scientific name Felis thinobius. Later he considered it a sand cat subspecies, which to date is widely recognised.
- F. m. meinertzhageni proposed by Pocock in 1938 was a sand cat skin from the Algerian Sahara.
- F. m. aïrensis proposed by Pocock in 1938 was a female specimen collected in the Aïr Mountains in 1937.
- F. m. scheffeli proposed by German zoologist Helmut Hemmer in 1974 was described on the basis of seven sand cats that had been captured alive in Pakistan's Nushki desert.
- F. m. harrisoni proposed by Hemmer, Grubb and Groves in 1976 was described on the basis of a skin and skull of an adult male sand cat captured in 1967 in Umm al Samim, Oman.
In 1974, F. m. margarita, F. m. thinobia and F. m. scheffeli were temporarily recognized as valid taxa. At the time, it was considered possible that sand cats eventually recorded in Afghanistan and Iran might constitute distinct subspecies. In 2005, F. m. margarita, F. m. thinobia, F. m. scheffeli and F. m. harrisoni were recognised as valid by W. Chris Wozencraft in the Mammal Species of the World, but F. m. meinertzhageni and F. m. aïrensis were considered synonyms of F. m. margarita. The Cat Classification Task Force of the Cat Specialist Group reviewed the existing information and in 2017 recognized only two subspecies, namely:
- F. m. margarita Loche, 1858: Morphologically distinguished by its smaller size and more yellow-colored spotted or striped fur; occurs in North Africa.
- F. m. thinobia (Ognev, 1927): Larger in size with reduced markings and a darker, greyer coat; occurs in West and Central Asia.
Phylogenetic analysis of tissue samples from all Felidae species revealed that the sand cat is part of an evolutionary lineage that is estimated to have genetically diverged between 6.7 and 6.2 million years ago. The sand cat probably diverged from the common ancestor of Felis species between 3.67 and 1.72 million years ago. The following cladogram shows the phylogenetic relationships of the sand cat as derived through analysis of nDNA:
The sand cat's fur is of a pale, sandy, light brownish-yellow color. Markings vary between individuals: some have neither spots nor stripes, some are faintly spotted, some have both spots and stripes. There are dark brown to blackish bars on the limbs, and the tail has a black tip with two or three dark rings alternating with buff bands. The head is sandy brown, whereas the lower and upper lips, chin, throat, and belly are white. Some individuals have a yellowish throat. The large, greenish-yellow eyes are ringed with white, and the nose is blackish. The lower part of the face is whitish, and a faint reddish line runs from the outer corner of each eye across the cheeks. The cat's whiskers are white and up to 8 cm (3.1 in) long. The sand cat is a small cat, characterized by a flat, wide head, short legs, and a relatively long tail of 23–31 cm (9.1–12.2 in). It stands 24–36 cm (9.4–14.2 in) at the shoulder and weighs 1.5–3.4 kg (3.3–7.5 lb). The head-and-body length ranges from 39–52 cm (15–20 in). The 5–7 cm (2.0–2.8 in) long ears are set low, giving a broad, flat appearance to the head. The ears are tawny at the base and tipped with black, and more pointed than those of the Pallas's cat (Otocolobus manul).
In Central Asia, the sand cat's winter coat is very long and thick, with hairs reaching up to 2 in (5.1 cm) in length. The sand cat's claws on the forelimbs are short and very sharp, and claws on the hind feet are small and blunt. The undersides of its paws are protected from extreme temperatures by a thick covering of fur. The long hairs growing between its toes create a cushion of fur over the foot pads, helping to insulate them while moving over hot sand. This feature makes the cat's tracks obscure and difficult to identify and follow.
Its skull is arched in lateral outline with wide zygomatic arches. The pinnae of the ears are triangular, and the ear canal is very wide, giving the cat an enhanced sense of hearing. The auditory bullae and the passages from the external ears to the ear drums are greatly enlarged compared to other small wild cats; the inner parts of the ears are protected from foreign objects by long, closely spaced white hairs. The sand cat's outer ear is similar to that of a domestic cat, but its ear canal is about twice the size. The magnitude of acoustic input-admittance is about five times higher than of a domestic cat. Additionally, hearing sensitivity of the sand cat is about 8 decibels greater than that of the domestic cat. It has a bite force quotient at the canine tip of 136.7.
Distribution and habitat
The sand cat inhabits both sandy and stony deserts. It is widely though not contiguously distributed in the deserts of North Africa, Southwest and Central Asia. It prefers flat or undulating terrain with sparse vegetation, and avoids bare sand dunes, where little prey is available. It retreats into burrows when climatic conditions are extreme such as temperatures of −5 °C (23 °F) or 52 °C (126 °F).
In the Western Sahara, sand cats were sighted and photographed in the Dakhla-Oued Ed-Dahab region several times between 2005 and 2016. Sand cat kittens were sighted and photographed in this area in spring 2017 that were hidden beneath a tuft of Panicum turgidum grass. In Algeria, one individual was recorded near a salt cedar (Tamarix aphylla) mound in the Ahaggar Mountains in 2008. No confirmed records are known in Mauritania, Tunisia and Libya. In Mali's Lake Faguibine area, one individual was shortly sighted by night in 2011. In the Ténéré Desert, sand cats were observed in the 1980s and between 2008 and 2015. Sightings in Egypt's rocky Western and Eastern Deserts date to the mid 1980s. In the Sinai peninsula, sand cats were sighted in the mid 1990s.
On the Arabian Peninsula, sand cats were captured in Saudi Arabia's Mahazat as-Sayd Protected Area and encountered trapped in wire mesh fence surrounding the adjacent Saja/Umm Ar-Rimth Protected Area in the country's Najd region. In 2003, a sand cat was sighted in a gravel plain between dunes in the Al-Ain Region, Abu Dhabi. Several sand cats were recorded in a protected area in Abu Dhabi's western region between April and December 2015, after an absence of sightings for ten years.
In the late 1980s, four sand cats were radio-collared and tracked over a few months in southern Israel's Arabah Valley. In 1997, a sand cat was recorded in a Jordanian desert. In 2000 and 2001, sand cats were sighted and photographed by a camera-trap in a protected area near Palmyra in Syria.
In Iran, it occurs in arid flat plains and sandy desert of Abbas'abad Wildlife Reserve, Kavir National Park and Petergan Rural District. Between March 2014 and July 2016, sand cats were also observed at altitudes of 900–1,100 m (3,000–3,600 ft) in Sistan and Baluchestan Province, foremost in black saxaul (Haloxylon ammodendron) dominated habitat. In Pakistan, the first sand cat was detected in 1966 near the Lora River in Balochistan. In the late 1960s, sand cats were also encountered in the Chagai Hills, an extremely arid area comprising rolling sand dunes and stony plains at an altitude of about 1,200 m (3,900 ft).
In Central Asia, the sand cat was known to occur up to the late 1960s in the Karakum Desert from the Ustyurt Plateau in the northwest to the Kopet Dag Mountains in the south, and from the Kyzylkum Desert to the Syr Darya River and the northern border to Afghanistan. In spring 2013 and 2014, adult sand cats with kitten were photographed in the southern Kyzylkum Desert, indicating that the population is breeding.
Behaviour and ecology
The sand cat is a solitary cat except during the mating season and when a female has kittens. It communicates using scent and scratch marks on objects in its range and by urine spraying. It makes loud, high-pitched and short rasping sounds, especially when seeking a mate. Its vocalizations are similar to those of the domestic cat.
Its way of moving is distinct: with belly close to the ground, it moves at a fast run punctuated with occasional leaps. It is capable of sudden bursts of speed and can sprint at speeds of 30–40 km (19–25 mi) per hour. It buries its feces, covering it with sand. Four radio-collared sand cats in Israel moved long distances of 5–10 km (3.1–6.2 mi) in a single night. They were generally active throughout the night, hunting and travelling an average distance of 5.4 km (3.4 mi). They retired below ground at dawn and stayed in the burrow during the day. During the survey period, they used several burrows in their home ranges. Burrows are about 1.5 m (4.9 ft) deep and dug in slightly slanting ground with usually only a single entrance. Burrows with two or three entrances have also been observed. These burrows were either abandoned by foxes (Vulpes) or porcupines, or dug by gerbils or other rodents. In winter, sand cats stay in the sun during the day, but during the hot season, they are crepuscular and nocturnal.
A male sand cat in Israel had a home range of 16 km2 (6.2 sq mi). In Morocco, a male sand cat travelled 14.1 km (8.8 mi) in 30 hours. A female sand cat moved in an area of 13.4 km2 (5.2 sq mi) during six days, and two males had home ranges of 21.8 and 35.3 km2 (8.4 and 13.6 sq mi).
Hunting and diet
The sand cat preys foremost on small rodents, including lesser Egyptian gerbil (Gerbillus gerbillus), lesser Egyptian jerboa (Jaculus jaculus), and young of cape hare (Lepus capensis). Several individuals were also observed hunting greater hoopoe lark (Alaemon alaudipes), desert monitor (Varanus griseus), sandfish (Scincus scincus), horned viper (Cerastes cerastes) and sand viper (C. vipera). If they caught more than they could eat, they buried the remains for later consumption. They satisfied their moisture requirements from their prey but drank readily if water was available. Toubou people in the Ténéré Desert accounted of sand cats coming to their camps at night and drinking fresh milk. In Israel, remains of Egyptian spiny-tailed lizards (Uromastyx aegyptia) were found near burrows used by sand cats. They were observed preying on jirds (Meriones), Cairo spiny mouse (Acomys cahirinus), desert lark (Ammomanes deserti), fringe-toed lizards (Acanthodactylus) and short-fingered geckos (Stenodactylus).
Sand cats were collected in eastern Karakum Desert in the late 1950s. Their faeces and stomachs contained remains of tolai hare (Lepus tolai), yellow ground squirrel (Spermophilus fulvus), great gerbil (Rhombomys opimus), midday gerbil (Meriones meridianus), northern three-toed jerboa (Dipus sagitta), comb-toed jerboa (Paradipus ctenodactylus), great spotted woodpecker (Denorocopos major), collared dove (Streptopelia), Pander's ground jay (Podoces panderi), hoopoe (Upupa epops), crested lark (Galerida cristata), desert sparrow (Passer simplex), spotted rat snake (Spalerosophis diadema), Karelin's snake (Coluber karelini), wonder gecko (Teratoscincus), Tenebrionidae beetles, scorpiones, Phalangiidae and arthropods. In March 2018, a sand cat was recorded feeding on an Asian Houbara Chlamydotis macqueenii in the Kyzylkum Desert.
Oestrus in female sand cats lasts from five to six days, during which they frequently call and scent mark. After a gestation of 59 to 66 days, they give birth to a litter of two to three kittens. They weigh 39 to 80 g (1.4 to 2.8 oz) at birth, and have spotted pale yellow or reddish fur. They grow relatively rapidly, reaching three quarters of the adult size within five months. They are fully independent by the end of their first year and reach sexual maturity not long after. In some areas, sand cats give birth to two litters per year.
Of 228 sand cats born in zoos globally by 2007, only 61% of the kittens lived to day 30. They died primarily due to maternal neglect by first-time mothers. They can live up to 13 years in captivity. The life expectancy of wild sand cats has not been documented.
Habitat degradation is the major threat to the sand cat. Vulnerable arid ecosystems are being rapidly degraded by human settlement and activity, especially livestock grazing. The sand cat's small-mammal prey-base depends on having adequate vegetation, which may experience large fluctuations due to drought or declines due to desertification and loss of natural vegetation. They also may be killed in traps laid out by inhabitants of oases targeting foxes and jackals or in retaliation for killing their chickens. Occasional reports of animals shot in southeast Arabia have been made.
Felis margarita is listed on CITES Appendix II. Hunting is prohibited in Algeria, Iran, Israel, Kazakhstan, Mauritania, Niger, Pakistan, and Tunisia. No legal protection exists in Egypt, Mali, Morocco, Oman, Saudi Arabia, and the United Arab Emirates. Previously having been classified as near threatened, it has been downlisted to least concern in 2016, as the estimated size of the global population exceeds the threshold for a threatened category; the extent of decline of the global population is unknown.
The Jerusalem Biblical Zoo started a sand cat reintroduction project in Israel's Arava Desert. Several captive-born individuals from the zoo's population were kept in an acclimatization enclosure, but did not survive subsequent release into the wild.
Captive sand cats are highly sensitive to respiratory diseases and infection of the upper respiratory tract. This is the main cause of death in adults. The most common disease is infectious rhinotracheitis. With sand cats being very susceptible to respiratory infections, they have to be kept in very arid enclosures, where humidity and temperature do not fluctuate.
The captive population kept in the European Endangered Species Programme is offspring of 18 founders that originated almost exclusively on the Arabian Peninsula. Until December 2009, the global captive population comprised 200 individuals in 45 institutions, including 23 European zoos with 102 individuals. The captive population within the Species Survival Plan for sand cat is based on eight founders.
In January 2010, the Al Ain Zoo announced the first success of an in vitro fertilisation and embryo transfer procedure on sand cats, resulting in the birth of two kittens at its facilities. In July 2012, four sand cat kittens were born at the Ramat Gan Zoo as part of the European Endangered Species Programme.
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